Brazilian wetlands: their definition, delineation, and classi cation … · 2018-10-12 ·...

18
Brazilian wetlands: their denition, delineation, and classication for research, sustainable management, and protection W. J. JUNK a , M. T. F. PIEDADE b , R. LOURIVAL c , F. WITTMANN d, * , P. KANDUS e , L. D. LACERDA f , R. L. BOZELLI g , F. A. ESTEVES h , C. NUNES DA CUNHA i , L. MALTCHIK j , J. SCHÖNGART d , Y. SCHAEFFER-NOVELLI k and A. A. AGOSTINHO l a Instituto Nacional de Ciência e Tecnologia em Áreas Úmidas, Universidade Federal de Mato Grosso, Cuiabá, Brazil b Instituto Nacional de Pesquisas da Amazônia, Grupo MAUA, Manaus, Brazil c Ministério da Ciência, Tecnologia e Inovação Brasil, Ecology Centre - University of Queensland, Australia d Max Planck Institute for Chemistry, Biogeochemistry Department, Mainz, Germany e Laboratorio de Ecología, Teledetección y Eco-Informática, Instituto de Investigaciones e Ingeniería Ambiental, Universidad Nacional de General San Martín, Argentina f Instituto de Ciências do Mar, Universidade Federal do Ceará, Fortaleza, Brazil g Universidade Federal do Rio de Janeiro, Instituto de Biologia, Departamento de Ecologia, Laboratório de Limnologia, Rio de Janeiro, Brazil h Universidade Federal do Rio de Janeiro, Núcleo em Ecologia e Desenvolvimento Socio-Ambiental de Macaé, Brazil i Departamento Botânica e Ecologia, Universidade Federal de Mato Grosso, Cuiabá, Brazil j Lab. Ecologia e Conservação de Ecossistemas Aquáticos, Unisinos, Porto Alegre, Brazil k Instituto Oceanográco, Universidade de São Paulo, Brazil l Universidade Estadual de Maringá Nupélia, Maringá, Brazil ABSTRACT 1. Although 20% of Brazilian territory is covered by wetlands, wetland inventories are still incomplete. In 1993, Brazil signed the Ramsar Convention but a coherent national policy for the sustainable management and protection of wetlands has yet to be established. 2. Major gaps in the denition of a specic wetland policy are twofold: (1) the lack of standardized criteria by which wetlands are dened and delineated that reects the specic ecological conditions of the country and (2) the lack of a national classication of wetlands that takes into account specic hydrological conditions and respective plant communities. 3. In recent years, efforts have been made at a regional level to improve public awareness of the ecology of Brazilian wetlands, their benets to society, and the major threats endangering them. Studies have shown that wetlands play a crucial role in the regional hydrological cycle and provide multiple benets for local populations. Furthermore, Brazilian wetlands contribute signicantly to South American biodiversity. Therefore, wetland conservation and sustainable management should be given high legislative priority. 4. This article provides a synthesis of the current body of knowledge on the distribution, hydrology, and vegetation cover of Brazilian wetlands. Their denition, delineation, and classication at the national level are proposed in order to establish a scientic basis for discussions on a national wetland policy that mandates the sustainable management of Brazils extremely diverse and complex wetlands. This goal is particularly urgent in the face of the continuing and *Correspondence to: Florian Wittmann, Max Planck Institute for Chemistry, Dep. Biogeochemistry, Hahn-Meiner Weg 1, 55128 Mainz Germany. Email: [email protected]; [email protected] Copyright # 2013 John Wiley & Sons, Ltd. AQUATIC CONSERVATION: MARINE AND FRESHWATER ECOSYSTEMS Aquatic Conserv: Mar. Freshw. Ecosyst. 24:522 (2014) Published online 15 August 2013 in Wiley Online Library (wileyonlinelibrary.com). DOI: 10.1002/aqc.2386

Transcript of Brazilian wetlands: their definition, delineation, and classi cation … · 2018-10-12 ·...

Page 1: Brazilian wetlands: their definition, delineation, and classi cation … · 2018-10-12 · Brazilian wetlands, their benefits to society, and the major threats endangering them.

Brazilian wetlands: their definition, delineation, and classificationfor research, sustainable management, and protection

W. J. JUNKa, M. T. F. PIEDADEb, R. LOURIVALc, F. WITTMANNd,*, P. KANDUSe, L. D. LACERDAf,R. L. BOZELLIg, F. A. ESTEVESh, C. NUNES DA CUNHAi, L. MALTCHIKj, J. SCHÖNGARTd,

Y. SCHAEFFER-NOVELLIk and A. A. AGOSTINHOl

aInstituto Nacional de Ciência e Tecnologia em Áreas Úmidas, Universidade Federal de Mato Grosso, Cuiabá, BrazilbInstituto Nacional de Pesquisas da Amazônia, Grupo MAUA, Manaus, Brazil

cMinistério da Ciência, Tecnologia e Inovação – Brasil, Ecology Centre - University of Queensland, AustraliadMax Planck Institute for Chemistry, Biogeochemistry Department, Mainz, Germany

eLaboratorio de Ecología, Teledetección y Eco-Informática, Instituto de Investigaciones e Ingeniería Ambiental, Universidad Nacionalde General San Martín, Argentina

fInstituto de Ciências do Mar, Universidade Federal do Ceará, Fortaleza, BrazilgUniversidade Federal do Rio de Janeiro, Instituto de Biologia, Departamento de Ecologia, Laboratório de Limnologia, Rio de Janeiro, Brazil

hUniversidade Federal do Rio de Janeiro, Núcleo em Ecologia e Desenvolvimento Socio-Ambiental de Macaé, BraziliDepartamento Botânica e Ecologia, Universidade Federal de Mato Grosso, Cuiabá, BraziljLab. Ecologia e Conservação de Ecossistemas Aquáticos, Unisinos, Porto Alegre, Brazil

kInstituto Oceanográfico, Universidade de São Paulo, BrazillUniversidade Estadual de Maringá – Nupélia, Maringá, Brazil

ABSTRACT

1. Although 20% of Brazilian territory is covered by wetlands, wetland inventories are still incomplete. In 1993,Brazil signed the Ramsar Convention but a coherent national policy for the sustainable management andprotection of wetlands has yet to be established.

2. Major gaps in the definition of a specific wetland policy are twofold: (1) the lack of standardized criteria bywhich wetlands are defined and delineated that reflects the specific ecological conditions of the country and (2)the lack of a national classification of wetlands that takes into account specific hydrological conditions andrespective plant communities.

3. In recent years, efforts have been made at a regional level to improve public awareness of the ecology ofBrazilian wetlands, their benefits to society, and the major threats endangering them. Studies have shown thatwetlands play a crucial role in the regional hydrological cycle and provide multiple benefits for localpopulations. Furthermore, Brazilian wetlands contribute significantly to South American biodiversity.Therefore, wetland conservation and sustainable management should be given high legislative priority.

4. This article provides a synthesis of the current body of knowledge on the distribution, hydrology, and vegetationcover of Brazilian wetlands. Their definition, delineation, and classification at the national level are proposed in orderto establish a scientific basis for discussions on a national wetland policy that mandates the sustainablemanagement ofBrazil’s extremely diverse and complex wetlands. This goal is particularly urgent in the face of the continuing and

*Correspondence to: Florian Wittmann, Max Planck Institute for Chemistry, Dep. Biogeochemistry, Hahn-Meiner Weg 1, 55128 Mainz – Germany.Email: [email protected]; [email protected]

Copyright # 2013 John Wiley & Sons, Ltd.

AQUATIC CONSERVATION: MARINE AND FRESHWATER ECOSYSTEMS

Aquatic Conserv: Mar. Freshw. Ecosyst. 24: 5–22 (2014)

Published online 15 August 2013 in Wiley Online Library(wileyonlinelibrary.com). DOI: 10.1002/aqc.2386

Page 2: Brazilian wetlands: their definition, delineation, and classi cation … · 2018-10-12 · Brazilian wetlands, their benefits to society, and the major threats endangering them.

dramatic deterioration of wetlands resulting from large-scale agro-industrial expansion, and hydroelectric projects aswell as the projected impact of global climate change on hydrological cycles.Copyright # 2013 John Wiley & Sons, Ltd.

Received 05 February 2013; Revised 28 June 2013; Accepted 29 June 2013

KEY WORDS: Amazonian wetlands; cerrado wetlands; coastal wetlands; flood pulse; Pantanal; Ramsar Convention; wetlandvegetation; wetland policy

INTRODUCTION

Wetlands are among the most threatened ecosystemsworldwide despite several international treaties thatrecommend both their regular inventory and effortsaimed at their protection (Millennium EcosystemAssessment, 2005; Darwall et al., 2008; SCBD, 2010).Brazil is the fifth largest country in the world,covering an area in the Neotropics of about 8.5million km2, in which a wide variety of wetland typesoccupy an estimated 20% of the national territory(Junk et al., 2011). Different types of forestedwetlands cover about 30% of the humid tropics of theAmazon lowlands. This percentage decreases towardthe dryer areas in the northern and southern parts ofBrazil, but even in the savanna belts there areextensive wetland systems. Some of these cover tensof thousands of square kilometres, e.g. the PanatanalMatogrossense and the flooded savannas of theAraguaia River, including Bananal Island. Along theAtlantic coast, mangroves are found down to 28–30oS,covering about 13800km2 (Kjerfve and Lacerda,1993). Indeed, extended lagoons and connectedwetlands are characteristic of the entire Brazilian coast.

Wetlands provide many services for society, suchas water storage, the buffering of river and streamdischarge, groundwater recharge, sedimentretention, water purification, microclimate regulation,recreation and ecotourism, organic carbon storage,timber production, and the provision of non-timberproducts, medicinal plants, fish, agricultural products,drinking water for humans and livestock, and pastureland for animal husbandry. Furthermore theycontribute to cultural safeguarding by providinghome for traditional communities (MillenniumEcosystem Assessment, 2005).

Wetlands also contribute significantly to biodiversity(Gopal et al., 2000). Predictable flood-pulsing wetlandscan be considered as centres of speciation, as evidenced

by the many endemic species of terrestrial invertebratesand trees in Amazonian floodplains (Erwin and Adis,1982; Adis, 1997; Junk, 2000; Wittmann et al., 2013,and the development of morphological, anatomicaland physiological adaptations as well as specificlife-history traits of invertebrates (Adis and Junk,2002). Furthermore, wetlands influence in multipleways species diversity of adjacent upland and deep-water habitats. For example, in the Pantanal, thereare 104 wetland-dependent bird species and 286upland species. Some upland species are, outside thePantanal, in danger of extinction, such as thehyacinth macaw (Anodorhynchus hyacinthinus), buthave large populations inside the Pantanal (Junk et al.,2006). Marine fish species, such as Mugil spp.,Anchoa spp., Centropomus spp., Sphoeroides spp.,and Lutjanus spp., use the mangroves for spawningand as juvenile nurseries, analogous to the use byriverine freshwater fish species of the adjacentfloodplains. The long-distance spawning migrationsof some characids, such as Semaprochilodus spp.,Prochilodus nigricans, and Brycon melanopterus,link deep-water habitats of the river channel to theperiodically inundated floodplains (Junk, 2007).The negative impacts of human-imposed pulseregulations on biodiversity and fisheries have beendemonstrated in several studies of the Paraná Riverand its floodplain (Hoeinghaus et al., 2009; Barlettaet al., 2010).

The mean global value of these services wasoriginally estimated by Costanza et al. (1997),while more recent estimates were based on effortsof an international initiative on ‘The Economics ofEcosystems and Biodiversity’ (TEEB, 2013). Bothestimates show values for wetlands that are higherthan those of most other ecosystems; however,economic valuations of specific wetlands may varywidely. For example, according to Seidl and

W. J. JUNK ET AL.6

Copyright # 2013 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 24: 5–22 (2014)

Page 3: Brazilian wetlands: their definition, delineation, and classi cation … · 2018-10-12 · Brazilian wetlands, their benefits to society, and the major threats endangering them.

Moraes (2000), who used methods similar to thoseof Costanza et al. (1997), the global wetland valuefor services of the Pantanal of Nhecolândia is halfthat of the annual 14 785 US$ ha-1 estimated bythe latter authors. Regardless of exact values,available data already point out the magnitude ofthe economic, ecological, and social value thatintact wetlands provide to Brazilian society.

Despite their geographic extension, diversity, andeconomic importance, wetlands are rarely mentionedin federal legislation, state constitutions, orenvironmental legislation. There is no national policythat regulates their protection and management.Only the Pantanal is distinguished as a NationalHeritage site by the 1988 constitution. Moreover, asnoted above, some large wetlands, e.g. thefloodplains of the Amazon River and several of itstributaries, as well as those of the Paraná and ParaguaiRivers, along with the Pantanal Matogrossense,cross national boundaries into other countries, suchthat international efforts are needed if harmoniousenvironmental policies are to be established.

In 1993, Brazil signed the Ramsar Conventionand since then has declared several Ramsar sites,but it has been very slow in the implementation ofwetland inventories and in wetland classification(Diegues, 1994, 2002). This process is complicatedby the dozens of local terms for different wetlandtypes and by the lack of broadly acceptedparameters to define wetlands, both of which areessential preconditions for a modern and efficientnational wetland policy aligned with the terms ofthe Convention. Such a policy is essential,particularly considering the impact of recurrentcatastrophic floods and droughts throughout thecountry’s territory and the likelihood that suchevents will increase in response to the forecastedglobal changes in climate (IPCC, 2007).

Recently, a few Brazilian institutions have initiatedefforts to achieve wetland classification. Theseinstitutions include the National Institute for Scienceand Technology in Wetlands (INCT-INAU) at theFederal University of Mato Grosso, Cuiabá, theworking group Monitoring Amazonian Wetlands(MAUA) at the National Amazon Research Institute(INPA) at Manaus, the Laboratory for Ecology andConservation of Aquatic Ecosystems (UNISINOS) atSão Leopoldo, Rio Grande do Sul, the Center for

Research in Limnology, Ichthyology and Aquaculture,at the State University of Maringá, Paraná, theInstitute of Marine Sciences of the FederalUniversity of Ceará (LABOMAR-UFC), the Instituteof Oceanography of Sao Paulo University, theInstitute of Oceanography of the Federal Universityof Rio Grande, and the Museu Paraense EmilioGoeldi in Belém, Pará. Consequently, classificationsystems are already available for Brazilian mangroves(Kjerfve and Lacerda, 1993), the permanent swampsof the cerrado (veredas) (Ribeiro and Walter, 1998;Araújo et al., 2002), parts of the semi-arid north east(Maltchik et al., 1999), the southern part of thecountry (Maltchik et al., 2003, 2004), the upperParaná River floodplain (Thomaz et al., 2004), thePantanal (Nunes da Cunha and Junk, 2011a), thewetlands of the Amazon basin (Junk et al., 2011),and the wetland habitats of the central AmazonRiver floodplain (várzea) (Junk et al., 2012b).

In this paper, a definition of Brazilian wetlands isproposed that corresponds to the specific hydrologicalconditions of the country and suggests a basisfor wetland delineation. A short ecologicalcharacterization of major wetland types is presentedand a hierarchical wetland classification method isintroduced that considers hydrological and vegetationparameters as the main wetland attributes. Theclassification is discussed in the context of othernational and international classification efforts. Asthe first step in the formulation of a Brazilian wetlandpolicy, it aims to contribute to worldwide efforts toachieve a better understanding of the extent,structures, and functions of wetlands as well as thethreats that endanger them.

PRECIPITATION, RIVER HYDROLOGY, ANDTHEIR IMPACT ON FLOOD PULSES IN

BRAZILIAN WETLANDS

Precipitation is not uniformly distributed within thedifferent regions of Brazil; rather, rainfall is highestin the north west (>3500mmyr-1) and lowest in thesemi-arid north east (300mmyr-1). Moreover, mostof the country’s regions face pronounced dry andwet seasons, with the exception of the rainfall-richnorth-western tropical forests and the southerncoastal areas (Figure 1).

BRAZILIAN WETLANDS 7

Copyright # 2013 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 24: 5–22 (2014)

Page 4: Brazilian wetlands: their definition, delineation, and classi cation … · 2018-10-12 · Brazilian wetlands, their benefits to society, and the major threats endangering them.

Large fluctuations in water level, also called‘flood-pulsing’, differentiate most wetlands in Braziland other countries of the tropics and sub-tropicsfrom those in countries at higher latitudes.Wetlands in the latter include bogs, fens, and mires,which typically have a relatively stable water level.In regions exposed to flood-pulsing, excessprecipitation during the rainy season leads toperiodic sheet-flooding of large, flat interfluvialareas, the periodic filling-in of depressions withwater, and the lateral inundation of large areasalong streams and rivers. These events result in theformation of extended river floodplains along mostof the large rivers and a dense network of riparianwetlands along streams and low-order rivers. Floodpulses are monomodal and predictable in the largefloodplains because they are part of the cycle of wetand dry seasons at regional or even continentalscale. The average amplitude of large-river floodpulses can exceed 10m in Amazonia but areconsiderably lower towards the south, depending onchanges both in the total amount and theperiodicity of the rainfall (Figure 2). Pulses ininterfluvial wetlands reach up to 2m on average.Lower-order rivers are subjected to short, unpredictable

flood pulses of varying height, according to theintensity of local rainstorms (Figure 3), whiledepressions in the semi-arid north east haveunpredictable inundations of short duration everyfew years. Structures and functions of large riverfloodplains are described by the flood pulseconcept (Junk et al., 1989; Junk and Wantzen,2004; Junk, 2005).

Coastal wetlands with direct marine influence aresubject to plurimodal, predictable tidal pulseswhereas, further inland, wetlands in coastal sandplains are subject to short, unpredictable, ormonomodal pulses during the rainy season. A generalclassification of the flood pulses is given in Table 1.Only a few Brazilian wetlands are permanently wetand have a mostly stable water level.

CHARACTERISTICS OF BRAZILIANWETLANDS AND PROPOSALS FOR THEIR

DEFINITION AND DELINEATIONThe extended terrestrial phases in flood-pulsingwetlands lead to the occupation of higher-lyingwetland areas by a specific vegetation comprisingwoody and herbaceous plants with a large ecologicaltolerance of flood and drought stress. In the Amazonrainforest, highly flood-tolerant, species-richfloodplain forests dominate wetlands. In savannaareas, the severe drought stress that oftencharacterizes the terrestrial phase favours a mixtureof savanna vegetation and forested patches(Drechsler et al., 2009; Lourival et al., 2011; Nunesda Cunha and Junk, 2011b). Wild fires are additionalpotential stress factors for wetland biota. Aquatic andpalustrine herbaceous plant species and invertebratesusually develop during the aquatic phase, often fromseed banks in the sediments or from vegetative restingstages. Fishes, aquatic birds, insects, reptiles,amphibians, and mammals tend to recolonizetemporary wetlands by emigration from permanentlyaquatic habitats. Then, at the beginning of the nextdry period they return to the permanent water bodiesor become stranded, die, and are incorporated withinterrestrial food webs.

Some wetlands, such as the large-riverfloodplains and the large periodically inundatedsavannas, cover huge areas and are very complex.For instance, the main-stem Amazon River

Figure 1. Precipitation curves for different areas of Brazil (Salati andMarques, 1984, completed by J. Schöngart).

W. J. JUNK ET AL.8

Copyright # 2013 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 24: 5–22 (2014)

Page 5: Brazilian wetlands: their definition, delineation, and classi cation … · 2018-10-12 · Brazilian wetlands, their benefits to society, and the major threats endangering them.

floodplain (várzea) covers 98 110 km2 (Melack andHess, 2010); the periodically flooded savannas inRoraima in Northern Amazonia (and Rupununiin Guiana), about 16 500 km2 (Melack and Hess,2010); the periodically flooded savannas of the

Guapore River, extending into the Boliviansavannas of the Mamoré and Mortes Rivers(Llanos dos Moxos), 92 100 km2 (Hamilton et al.,2004); the Pantanal at the border of Brazil,Bolivia, and Paraguay, 109 590 km2 (Hamilton

Figure 2. Flood curves for large Brazilian rivers. The geographic positions of the respective data collection points are indicated by the numbers on themap. The curves represent the water-level fluctuation (m) during the annual cycle between 1970 and 2010. The black curve represents the daily meanwater level and the curves above and below it the daily maximum and minimum values for this period (for comparative reasons, the absolute minimumvalue of all curves was defined as zero on the height scale).The numbers below the river names indicate the mean values of the flood pulse

amplitude (m).

BRAZILIAN WETLANDS 9

Copyright # 2013 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 24: 5–22 (2014)

Page 6: Brazilian wetlands: their definition, delineation, and classi cation … · 2018-10-12 · Brazilian wetlands, their benefits to society, and the major threats endangering them.

et al., 1996); and the Araguaia River wetlands,including Bananal Island, 58 600 km2 (Melack andHess, 2010). Also of note are the coastal tidalwetlands, mostly mangroves and salt marshesalong the Maranhão/Pará littoral, which coverabout 7000 km2 (Lacerda, 2001) (Figure 4).

Many of these wetlands include elevated andpermanently dry areas of up to a few squarekilometres, deriving from outcropping base-rocks(inselbergs), palaeo-fluvial terraces, or ancient sea-level relict sand ridges. In shallow flooded savannaareas, termite mounds form small, permanentlydry islands of a few square metres each

(hyperseasonal termite savannas; Eiten, 1983).These permanently dry islands are of utmostimportance as periodic refuges of terrestrialorganisms and they contribute decisively to themaintenance of biodiversity cycles, functions, andprocesses. They must therefore be considered asindispensable parts of the large wetland systems.The extraordinary habitat diversity of the largeNeotropical wetlands requires individual habitatclassifications, as already proposed for thePantanal (Nunes da Cunha and Junk, 2011a)and the central Amazon River floodplain (Junket al., 2012b).

Figure 3. Daily precipitation and discharge in a low-order savanna stream near Cuiaba (according to Wantzen, 2003).

Table 1. Types of flood pulses and affected wetlands

Predictability Frequency Amplitude Wetland type

Predictable Monomodal High Large river floodplainsLow Large interfluvial wetlands, wetlands in coastal sand plains

(e.g. the Lençóis maranhenses)Predictable Polymodal Varying Tidal wetlandsUnpredictable Polymodal Varying Wetlands along low-order rivers, in depressions, and in

coastal sand plainsUnpredictable Pluriannual Low Wetlands in semi-arid zones of north-eastern Brazil

W. J. JUNK ET AL.10

Copyright # 2013 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 24: 5–22 (2014)

Page 7: Brazilian wetlands: their definition, delineation, and classi cation … · 2018-10-12 · Brazilian wetlands, their benefits to society, and the major threats endangering them.

Despite their smaller sizes, wetlands in the drynorth east, along the tropical coast to the south eastand in the wet subtropical portion of Brazil, play asignificant role in the regulation of the regionalhydrological regime and maintenance of biodiversity.Many of these wetlands harbour endemic andendangered species (Maltchik et al., 1999, 2003,2004), and provide water and food for many localcommunities and for livestock. They also serve asessential sources of recreation (Esteves, 2011).

Traditional communities have developed manystrategies to use specific wetland resources duringthe terrestrial and aquatic phases. Nevertheless,politicians and urban planners often considerwetlands as wastelands, allowing their exploitationas solid-waste dumping sites or housing areas ortheir disruption by road construction. In manyregions modern agro-industries have destroyedwetlands, and thus their multiple benefits, by

converting large areas of wetlands into croplands.In southern Brazil, this transformation is alreadyextensive. The resulting reduction in wetland buffercapacity together with the increased surface run-offleads to annual catastrophic floods, as occurred in2011 in Minas Gerais, with many victims and heavylosses of public and private goods (Junk et al., 2012a).Furthermore, the leaching of fertilizers and pesticidesas well as the run-off of untreated sewage from urbanareas causes a deterioration of water quality in rivers,streams and other downslope wetlands. Coastalwetlands have been destroyed by the expansion ofcities and in response to demands for waterfront orscenic homes. Brazil has enormous hydroelectricpotential but some of its reservoirs have destroyedriver floodplains, interrupting the longitudinalconnectivity of rivers and damaging downstreamwetlands through the alteration of flood pulses,sediment load, and other limnological parameters, as

Figure 4. Distribution of major wetlands in northern South America (according to Junk, 2007).

BRAZILIAN WETLANDS 11

Copyright # 2013 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 24: 5–22 (2014)

Page 8: Brazilian wetlands: their definition, delineation, and classi cation … · 2018-10-12 · Brazilian wetlands, their benefits to society, and the major threats endangering them.

shown for the Paraná River and its floodplain(Hoeinghaus et al., 2009; Barletta et al., 2010). Thesenumerous adverse effects highlight the importance ofincluding the flood-pulse concept in the definition ofBrazilian wetlands and integrating it into plans fortheir protection.

We propose the following definition and delineationof Brazilian wetlands:

Wetlands are ecosystems at the interface betweenaquatic and terrestrial environments; they may becontinental or coastal, natural or artificial,permanently or periodically inundated by shallowwater or consist of waterlogged soils. Their watersmay be fresh, or highly or mildly saline. Wetlands arehome to specific plant and animal communitiesadapted to their hydrological dynamics.

The extent of a wetland can be determined by the borderof the permanently flooded or waterlogged area, or inthe case of fluctuating water levels, by the limit of thearea influenced during the mean maximum flood. Theouter borders of wetlands are indicated by the absenceof hydromorphic soils and/or hydrophytes and/orspecific woody species that are able to grow inperiodically or permanently flooded or waterloggedsoils. The definition of a wetland area should include,if present, internal permanently dry areas as thesehabitats are of fundamental importance to themaintenance of the functional integrity andbiodiversity of the respective wetland.

PROS AND CONS OF SOME EXISTINGCLASSIFICATION SYSTEMS

The scientific literature contains many definitions ofwetlands as well as systems for their classification(summarized in Mitsch and Gosselink, 2008).However, most of them emphasize wetlands withpermanent or long-term wet conditions whilelargely neglecting those subjected to flood-pulsing,with long terrestrial phases. Organic matteraccumulation as a result of permanent shallowinundation or long-term waterlogging is a goodindicator of permanently wet conditions. In contrast,flood-pulsing wetlands with long terrestrial phases donot exhibit accumulation of organic matter asperiodic aeration facilitates the decomposition oforganic matter. Thus, by exclusively focusing on

hydric soil indicators we fail to achieve wetlanddelineation and protection, not only in Brazil but alsoworldwide, i.e. in all regions where there is apronounced seasonality in rainfall. In fact, all of thewell-known large African wetlands, such as theOkawango Delta, the Niger River floodplain, and theSudd, annually undergo an extended dry phase.Many Australian wetlands become wet only everyfew years but they are of utmost importance for themaintenance of biodiversity. Likewise, in temperateregions, large areas of river floodplains fallperiodically dry. In many cases, wetland status ofhigh-lying floodplain areas inundated only duringpeak floods is not recognized, often facilitatingexploitation of these areas as cropland andprotected by dikes, e.g. along the Mississippi,Missouri, and Ohio Rivers in the USA and theRhine River in Germany. This has had far-reachingadverse consequences for the flood regime, nutrientcycles, and habitat and species diversity not only offormer and still active river–floodplain complexesbut also of river deltas and the adjacent sea (Mitschand Day, 2006).

The Ramsar Convention (Scott and Jones, 1995)classifies wetlands worldwide, differentiating betweenmarine and coastal, inland, and man-made wetlandsystems. These systems are subdivided for hydrologicalcharacterization using the terms subtidal, intertidal,perennial, intermittent, permanent, and seasonal.Sub-units are characterized by geomorphological,hydrological, and/or botanical parameters. Theinclusion of shallow-water coral reefs in thedefinition is questionable and may over-extend thewetland concept. The classification system ofwetlands and deep water habitats of the US Forestand Wildlife Service (UFWS) (Cowardin et al.,1979) differentiates between marine, estuarine,riverine, lacustrine, and palustrine systems and usesthe terms subtidal, intertidal, tidal, lower perennial,higher perennial, intermittent, limnetic and littoralfor hydrological classification of the subsystems.Classes are characterized by geomorphological,hydrological, and/or botanical parameters. Thehydrogeomorphic classification of Brinson (1993)relies on geomorphic, physical, and chemicalparameters to provide a better understanding of therelationship between organisms and their environment.It is ‘a generic approach to classification and not a

W. J. JUNK ET AL.12

Copyright # 2013 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 24: 5–22 (2014)

Page 9: Brazilian wetlands: their definition, delineation, and classi cation … · 2018-10-12 · Brazilian wetlands, their benefits to society, and the major threats endangering them.

specific one to be used in practice’ (Brinson, 1993).Based on the Australian experience, Semeniuk andSemeniuk (1995) proposed a geomorphic approach,combining landforms and degrees of wetness, to theglobal classification of inland wetlands. The authorscorrectly extended the definition of the peripheralwetland boundary to periodic ‘dampness, or hydricsoils or vegetation indicative of wet conditions.’ Theirsystem does not reflect the geomorphic heterogeneityof large-river floodplains and internal deltas such asthe Amazonian large river floodplain, the Pantanal,and the Okawango Delta. There are two classificationsystems for Argentinian wetlands: Neiff (2001)differentiates between nine types, using 12 parametersto describe their geomorphology, soils, fire stress,vegetation, fauna, water origin, and severalhydrological factors; Brinson and Malvárez (2002)also differentiate between nine types, but useclimate, hydrology, soils and the regional vegetationas criteria.

The problems arising during the elaboration ofclassification systems were discussed by Finlaysonand Van der Valk (1995), who pointed out thenecessity of resolving differences between regionalwetland definitions and regional typologies. Theyalso drew attention to the need to standardize datacollection and disseminate new technologies inorder to establish ample international inventories.Indeed, many definitions and classification systemswere formulated decades ago for specific purposesand do not correspond to current scientific andregulatory requirements. Modern approaches, suchas the Asian Wetland Inventory (AWI), providepowerful tools for the Assessment and Monitoring ofWetland Biodiversity and Wise Use (Lopez et al.,2002). The need for a better wetland classification wasrecognized by the Ramsar Scientific and TechnicalReview Panel, which called for the ‘developmentand testing of a hydro-geomorphically-basedsystem of wetland classification’ (Davidson andFinlayson, 2007).

The new Brazilian classification differentiatesbetween coastal, inland, and artificial wetlandsand concentrates on hydrology and vegetationcover. Hydrology is the most important factordetermining wetland characteristics and is givenhighest priority. The approach is a practical onesince human impact on wetlands in Brazil and in

most other countries often starts with destructionof the natural vegetation cover, e.g. by timberextraction, cattle ranching, and crop plantations,which in turn cause changes in the hydrologicalregime through water abstraction, drainage, floodcontrol, and reservoir construction and is inevitablyfollowed by inappropriate civil construction. Thesesteps can be monitored by remote-sensing techniquesand the consequent measures required for wetlandprotection, including proposals for sustainablemanagement, can easily be explained to politicians,planners, decision-makers, and the public.Nonetheless, the large number of Brazilian wetlandswith oscillating water levels requires that greateremphasis be placed on the different types of floodpulses, which are under-represented in all of theclassification systems discussed above. While thehydro-geomorphic arguments provided by Brinson(1993) and Semeniuk and Semeniuk (1995) are veryhelpful from a scientific point of view, they contributelittle to the continuing political discussion on wetlandmanagement in Brazil.

For management purposes, the inclusion of localterms in national classification systems, as was doneby Gopal and Sah (1995) for Indian wetlands, islikely to be beneficial because it often increases thelocal population’s willingness to accept the imposedregulations for the sustainable management andprotection of wetlands. This approach is included inthe habitat classification systems provided for largeand complex wetland systems, such as the Pantanal(Da Silva et al., 2000; Nunes da Cunha and Junk,2011a) and the central Amazon River floodplain(Junk et al., 2012b).

PROPOSAL FOR THE CLASSIFICATION OFBRAZILIAN WETLANDS

The current classification for Brazilian wetlandsuses the structure proposed for the classification ofAmazonian inland wetlands (Junk et al., 2011). Itdiffers, however, in that one category of theregional Amazonian classification system, whichdifferentiates wetlands on the basis of water colour(white-water, black-water, and clear-water rivers),indicative of physicochemical conditions, has beenremoved from the national classification system,since it is applicable only to the Amazon and not

BRAZILIAN WETLANDS 13

Copyright # 2013 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 24: 5–22 (2014)

Page 10: Brazilian wetlands: their definition, delineation, and classi cation … · 2018-10-12 · Brazilian wetlands, their benefits to society, and the major threats endangering them.

to the entire country. However, in the future, thephysical and chemical characteristics of waters andsoils are likely to provide the basis for a moredetailed classification.

The Brazilian classification of wetlands issegregated into three levels: (1) systems, (2) unitsdefined by hydrological parameters, and (3) unitsdefined by higher plants, as shown in Figure 5. Thefirst (system) level is divided into three categories:

1. Coastal wetlands are defined as all wetlands,permanent or temporary, with fresh, brackish, orsaline waters, under direct influence of the tides, orsubject to saline intrusions, or influenced by theatmospheric deposition of dissolved or particulatesubstances and/or propagules from the ocean.

2. Inland wetlands are defined as all wetlands,permanent or temporary, with fresh, saline, or saltwater, that are located in the Brazilian inland andare thus without direct or indirect marine influence.

3. Artificial wetlands are all wetlands, coastal orinland, derived from human activities either inorganized (e.g. fish farms, rice paddy plantations),or unorganized forms (wetlands around reservoirsor those that progressively develop by thedamming of streams or that form in depressionscaused by the excavation of soil for roadconstruction, etc.).

The second hierarchic level is based on hydrologicalparameters and is composed of five subsystems: threedescribe coastal wetlands and two describe inlandwetlands. The two inland wetland subsystems aredivided into three orders and two sub-orders.The differentiation into sub-units emphasizes theimportance of hydrology and acknowledges thehydrological diversity of Brazilian natural inlandwetlands and wetland systems. This approach wasalso used in the classification of Amazonian inlandwetlands (Junk et al., 2011).

The hydrological characteristics of wetlandsidentify the origin of their waters (mainly fromrain, a parent river, or the sea) and whether theyare permanent, with a rather stable water level, orsubject to fluctuating (pulsing) water levels and thusto dry and wet periods. As noted above, themajority of Brazilian wetlands belong to thecategory of pulsing systems. These are classified withrespect to the amplitude, duration, predictability,and frequency of the flood pulse (Table 1).

The third hierarchic level is based on thecommunity structure and the occurrence of higherplants and is divided into classes, subclasses, andmacrohabitats. The communities of higher plants areparticularly appropriate for wetland classification(Drechsler et al., 2009), especially considering theirvisibility, dynamics, and longevity, all of whichrespond and reflect environmental conditions overperiods of months, years (herbaceous plants),decades, and even centuries (trees) (Naiman andDecamps, 1997; Casanova and Brock, 2000; Lourivalet al., 2011). At the macrohabitat level, the absenceof herbaceous plants can also serve as a criterion, forexample in descriptions of sandbanks, rocky shores,rocky outcrops, and steep erosion cliffs.

Current knowledge of the community structure ofwetland vegetation and the occurrence of speciesvaries between the different wetland systems.Species diversity is very large and cannot bediscussed here in detail. One of the challenges offuture studies is the characterization of macrohabitatsby species lists and the determination of specificindicator species. The following provides a briefsummary of the wetland vegetation in coastal andinland wetlands.

The vegetation of coastal wetlands is characterizedby mangroves, estuaries, and other types of wetlandvegetation communities. Mangrove species compositionis well documented for classification purposes(Schaeffer-Novelli et al., 1990; Bigarella, 2001;Menezes et al., 2008) but there is much lessinformation on the vegetation of the other coastalsubsystems, because of the high diversity of theseenvironments, ranging from temporary to permanentwetlands along a salinity gradient of freshwater tohypersaline (Araújo and Henriques, 1984; Irganget al., 1984; Irgang and Gastal, 1996; Costa andDias, 2001; Bove et al., 2003). In coastal lagoonsother salt-tolerant vegetation types may develop inaddition to mangroves. In particular, salt marsheswith Spartina spp., Salicornia spp., Juncus spp.,Paspalum spp., Crenea spp., Sesuvium spp., Cyperusspp., Batis spp., and Sporobolus spp are prevalent(Costa and Davy, 1992; Araújo et al., 1998).

Inland wetlands are very diverse with respect totheir hydrology and vegetation cover. The majorityof central Amazonian wetlands are forested(Schöngart et al., 2010; Wittmann et al., 2010;

W. J. JUNK ET AL.14

Copyright # 2013 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 24: 5–22 (2014)

Page 11: Brazilian wetlands: their definition, delineation, and classi cation … · 2018-10-12 · Brazilian wetlands, their benefits to society, and the major threats endangering them.

Veredas

Cyperus giganteus

Long-term flooded or water-logged grasslands

Mangroves

Flooded grasslands

Freshwater wetlands subjected to tidal pulses

Salt marshes

Monodominant herbaceous swampsTypha dominguensis

Turfeiras

Palm forests of Mauritia flexuosa

Mixed forests

Mangroves at sea shore

2 Inland wetlands

2.1. Wetlands with relatively stable water level

Forested swampy wetlands

Palm forests of Copernicia prunifera

Swamps with multi-species herbaceous vegetation

Hedychium coronarium

Temporary lakes and ponds

1.3. Wetlands separated from the sea with fluctuating water level

Periodically flooded forests

Long-term flooded or water-logged forests

Coastal wetlands without permanent connection to the sea

Freshwater lagoons

Lagoons with different levels of salinity

Flooded forests

Flooded grasslands1.2. Wetlands separated from the sea with little fluctuating water level

Coastal lagoons subjected to tidal pulses

Rocky shores

Sandy shoresMangroves in river mouth

Hypersaline areas

Saltwater wetlands

1.1. Wetlands influenced by predictable tidal flood pulses

1 Coastal wetlands

MacrohabitatSubclassClassSuborderOrderSubsystemSystem Order

2.2.2. Polymodal, unpredictable pulses of short durationRiparian wetlands along small rivers (1-5th order)Rain water fed wetlands in small depressions

2.2.3. Multiannual, unpredictable pulses of short durationRain water fed wetlands in depressions in the semi-arid North-East

Floodplain of Paraná River

Other high-amplitude pulsing floodplains

Floodplains along Amazonian black- and clear-water rivers (igapós)

Depressions along roads

Repression of streams by roads

Wetlands around hydroelectric reservoirs

Paddy rice plantations

Drainage channels

3 Artificial wetlandsAquaculture systems

Water reservoirs (e.g. açudes)

Amazonian hydromorphic edaphic savannas (campinas and campi-naranas)

Savanas of Roraima

Floodplains along Amazonian white-water rivers (várzeas)

Other hydromorphic climatic savannas

Wetlands Guaporé River

2.2.1.2. Low amplitude

Large interfluvial wetlands at the middle Negro River

Pantanal matogrossense Araguaia River wetlands, Ilha do Bananal

2.2.1. Monomodal, predictable pulse of long duration

2.2.1.1. High amplitude

2.2. Wetlands with fluctuating water level

)

Figure 5. Classification system of Brazilian wetlands according to hydrological and botanical parameters (woody and herbaceous plants). 1For theclassification of subclasses and macrohabitats see Junk et al. (2012b); 2for the classification of subclasses and macrohabitats see Nunes da Cunhaand Junk (2011a); 3classification of subclasses and macrohabitats in preparation by Junk et al.; 4classification of subclasses and macrohabitats not

yet available.

BRAZILIAN WETLANDS 15

Copyright # 2013 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 24: 5–22 (2014)

Page 12: Brazilian wetlands: their definition, delineation, and classi cation … · 2018-10-12 · Brazilian wetlands, their benefits to society, and the major threats endangering them.

Wittmann, 2013). In nutrient-rich white-water riverfloodplains (várzeas), extended communities of fast-growing, highly productive, emergent, and free-floating aquatic macrophytes develop at rising andhigh waters in front of the floodplain forest(Piedade et al., 1991; Junk and Piedade, 1997). Atlow water, these areas are occupied by terrestrial,mostly annual grasses, sedges, and herbs. Aquaticmacrophytes are restricted to free-floating mats andemergent species because high flood pulses createunfavourable light conditions near the bottom indeep water. In shallow waters, the floodplain forestcanopy absorbs too much light for herbaceous plantgrowth in the understorey. Consequently, there arefewer aquatic macrophyte species in Amazonianforested wetlands than in cerrado wetlands, e.g. thePantanal, and the wetlands of southern Brazil(Table 2). In black-water river floodplains (igapós),coloured humic substances in the water absorb lightand the nutrient status is extremely low, whichtogether limit the growth of free-floating andemergent aquatic macrophytes during the aquaticphase and that of terrestrial species during theterrestrial phase.

Both the genesis of Amazonian white-sandhabitats (campinas and campinaranas) and thecomposition of their vegetation are poorlyunderstood. Hydrological conditions can varyconsiderably among the different campina types.Some, or at least parts of them, are waterlogged or

flooded during the rainy season and should beconsidered as wetlands. Herbaceous vegetation isscarce because of the very low nutrient status andextreme drought stress that characterize the dryseason. Little is known about Amazonian swampand riparian forests along low-order rivers. In thediscussion about the Amazonian rainforest, theyare erroneously included in the category of uplandrainforests. Clearly, a list of wetland indicatorspecies for these areas must be elaborated. Palmforests of Mauritia flexuosa (buriti) are a specificand widespread wetland type. They range in areafrom 55 000 to >100 000 km2, forming extendedswamps in moist depressions in the forests andsavannas of tropical South America and oftenstoring considerable amounts of organic material(Kahn, 1991; Ruokolainen et al., 2001; Householderet al., 2012).

Cerrado wetlands include veredas and wetgrasslands (Ribeiro and Walter, 1998), gallery forests(Felfili, 1995), but also large and very complexwetlands, such as the Pantanal Matogrossense(Nunes da Cunha et al., 2007). Forested beltscomposed of flood-tolerant trees occur along riverchannels, around floodplain lakes, and in moistdepressions. Dryer areas are commonly occupied bytree species that are highly drought- and fire-tolerantbut also resistant to several months of shallowflooding or waterlogging. The diversity of aquaticmacrophytes is much larger here as both shallow,transparent water and the absence of dense forestfavour a rich submerged and floating flora (Table 2).

On southern Brazilian plateaux there arepermanently wet grasslands on organic deposits(turfeiras) that are heavily threatened byreclamation for agricultural purposes. Ephemeralwetlands in the north-eastern dry region of Brazilare small (± 1 ha for ponds) and are prone toshallow flooding every couple of years for a fewweeks only (Maltchik et al., 1999). They are veryimportant for the maintenance of regionalbiodiversity, such as anurans. In coastal areas,wax palm swamps (Copernicia prunifera) replacethe buriti palm swamps (Mauritia flexuosa).

Among the most threatened of Brazilianwetlands are those of the Atlantic forest, for whichlittle information exists. The original vegetationcover of this biome has been reduced to 11.7% by

Table 2. Taxonomic diversity of trees and herbaceous plants in theAmazon River floodplain (várzea) (Junk and Piedade, 1993;Wittmann et al., 2010; Wittmann, 2013), the Pantanal (Pott and Pott,2000; Junk et al., 2006), and the freshwater wetlands of southernBrazil (Rio Grande do Sul) (Rolon et al., 2010)

Amazonfloodplain Pantanal

South Brazilianwetlands

Woody plantsTotal >1000 750 179Terrestriala none 400 nonePalustrineb >1000 350 179c

Herbaceous plantsTotal 390 1150 280Terrestrial 340 900 no dataAquatic/palustrine 50 250 280Endemics 68 tree species none 1 herb. speciesd

aIn non-flooded areas inside the Pantanal.bIn periodically flooded or waterlogged areas.cMostly in riparian habitats of the campos sulinos (Wittmann,unpublished).dRegnellidium diphyllum.

W. J. JUNK ET AL.16

Copyright # 2013 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 24: 5–22 (2014)

Page 13: Brazilian wetlands: their definition, delineation, and classi cation … · 2018-10-12 · Brazilian wetlands, their benefits to society, and the major threats endangering them.

deforestation (Ribeiro et al., 2009). Wittmann(2012) differentiated inland wetlands of theAtlantic forest in riparian forests along streamsand interfluvial montane fens, bogs, andhygrophile forests and provided a synopsis of theexisting literature on forest inventories and themost common wetland tree species. The wealth ofregional terminologies attributed to forestssubjected to phreatic flooding suggests that a largevariety of forest types should be expected but alsothat further work is required to allow theircomparison with respect to their floristics, ecology,and biogeography (Scarano, 2006). Informationon herbaceous plant communities is scarce.

Southern freshwater wetlands belong mostly tothe subclass of ‘swampy wetlands covered byherbaceous plants’, which occupy relatively smallareas. An inventory of 260 wetlands of 0.15–10 harevealed a low number of flood-tolerant trees anda highly diversified herbaceous aquatic andpalustric flora (Table 2, Rolon et al., 2010).

In Brazil, the number of artificial wetlands aswell as their size and extent dramatically increasedduring the 20th century. The increase reflects theincreasing number of hydroelectric power plantsand drought prevention measures. The stronglyand irregularly fluctuating water levels of thesesystems expose large areas of land to periodic dryor waterlogged conditions. There is also a growingnumber of rice paddy plantations within thewetlands of southern Brazil, and some 1500 ha ofmangroves have been converted into shrimp pondson the Brazilian coast (2oS–20oS) (Maia et al.,2006). The cumulative environmental impact ofthese cultivation areas has yet to be determined butit is certainly large and provokes the liberation ofagrochemicals (Lacerda et al., 2006) and increaseddemand for fresh water that is abstracted fromneighbouring intact rivers and wetlands.

The new classification of Brazilian wetlands ispresented in Figure 5.

DISCUSSION AND CONCLUSIONS

Classification systems are often dependent on theobjective and on the chosen parameters. ForBrazilian wetlands, the primary objective is to

offer a hierarchical classification system, whichwould provide scientists working in Brazil with themeans to position their wetland studies withinbroader national and international contexts,thereby facilitating comparisons between differentwetlands, with the advantage of understanding the(dis)similarities among them. A second and perhapsmore important objective is to provide a scientificbasis for politicians and decision-makers toelaborate wetland-specific policies and legislation.

Better regulations are urgently needed given thatabout 20% of the Brazilian territory is covered bywetlands that are being lost at alarming rates ashuman activities expand into these areas. Theparameters and indicators for wetland definitionand delineation and the classification itself, aspresented in this article, will equip Brazil with animportant tool for structuring and controlling thespread of artificial wetlands, while making thesustainable use of natural wetland systems anenvironmental, social, and economic asset.

The differentiation of coastal, inland, and artificialwetlands and their hierarchical classification basedon hydrology and the community structure ofhigher plants make the proposed system compatiblewith other classification systems worldwide, e.g. thesystem developed for Indian wetlands (Gopal andSah, 1995). Most of the wetland types described inthe Ramsar or USFWS classification can also befound in the new Brazilian classification, albeitoften at another hierarchical level, which gives thema different importance in the overall wetlandhierarchy. A clear advantage of the newclassification is that it allows the introduction ofadditional units without the need to modify theentire concept. Considering the large size of Braziland the still precarious level of wetland inventories,the description of additional wetland units, at leastat the macrohabitat level, can be expected.

Indeed, the classification includes, at the classand subclass level, highly complex wetlands thatextend over tens of thousands of square kilometresand require additional habitat classifications forscientific research, management, and protection.Some of these classifications already exist, e.g. forthe Pantanal (Nunes da Cunha and Junk, 2011a)and the Amazon River floodplain (várzea) (Junket al., 2012b). Others are in preparation, e.g. for

BRAZILIAN WETLANDS 17

Copyright # 2013 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 24: 5–22 (2014)

Page 14: Brazilian wetlands: their definition, delineation, and classi cation … · 2018-10-12 · Brazilian wetlands, their benefits to society, and the major threats endangering them.

Amazonian black-water river floodplains (igapós),and Paraná, Araguaia, and Guaporé Riverfloodplains, and can be integrated without majordifficulty into the general classification presentedhere. The classification can also be further specified,e.g. by subdividing macrohabitats into smaller unitsaccording to water chemistry and/or soil parameters.

The emphasis given to hydrological classificationparameters takes into account the specific climaticsituation of Brazil. Different rainfall patterns leadto the formation of permanent wetlands but alsoto large periodic wetlands subject to flood pulsesdiffering in amplitude, duration, frequency, andpredictability. These hydrological parametersunderlie the structure and function of wetlandsand their biodiversity, and thus also determinethe possibilities of wetland management andconservation. The fact that most inland wetlandsare subject to extended dry periods implies the riskthat developers and politicians regard these areasas permanently dry habitats in which naturalflooding is not considered an inherent attribute ofthe system, but rather a catastrophic event thatmust be avoided or controlled. This view haspromoted the encroachment of conventionalagriculture and infrastructure development withinwetlands, as was emphasized during recentdisputes in the Brazilian Parliament over the newForest Code (Piedade et al., 2012).

According to Finlayson (2012) ‘the extensiveeffort to develop international policy, supportedby a substantial and expanding information base,has not stopped and reversed the global loss and/or degradation of wetlands’ because nationalgovernments have often failed to implement fully therecommendations of the Ramsar Convention. TheBrazilian National Report on the Implementation ofthe Ramsar Convention on Wetlands of 2012indicates in Section 3.1.1 the existence of acomprehensive wetland inventory (BRASIL, 2012).As discussed herein, there is extensive informationon Brazilian wetlands but it is far from beingcomprehensive. Under Point 1.3.1, the reportconfirms the existence of a national wetlands policybut in the additional information the authors statethat ‘there is no specific policy for wetlands. TheBrazilian Government believes that the best strategyfor the country is to enforce the existing extensive

environmental legislation, rather than creating a newpolicy instrument specifically focusing on wetlands.’However, the continuing political discussion on thenew Brazilian Forest Code shows that this strategyprovides very little protection for the country’swetlands (Piedade et al., 2012). This perspectivecreates serious impediments to the Braziliansustainability agenda and the country’s commitmentsto the Ramsar Convention while compromising theopportunities for the wise use of Brazilian wetlands.

Conservation is also a complex issue whendefinitions and delineation are not well structuredand clarified. Wetlands offer an extra level ofcomplexity for institutions in charge of planning theirprotection. The nature of the aquatic–terrestrialinterface of wetlands is a shifting one, spatially,seasonally and temporally, which must beappreciated by planners and decision-makers.Recent studies have provided several potentialdirections for future research aimed at developingplans for wetland protection, evolution, andpersistence (Beger et al., 2010). The importantcontribution of wetlands to water purification,flood control, and food production, both land- andwater-based, and the maintenance of high levels ofbiodiversity and the rich genetic repository of theirorganisms are just a few issues that clearlydistinguish wetlands in terms of their ecosystemrelevance to society. The second goal of theRamsar Convention priorities for the next fewyears is to propose the designation of new Ramsarsites, to ensure the representativeness of the varioustypes of wetlands in the respective countries.However, this implies the need both for aclassification system and a comprehensive inventoryof the wetlands.

Certainly, this article is only a first step towardsresolving the very complex scientific andadministrative problems related to Brazilian wetlands,but we hope that it will stimulate and ultimately helpto frame the scientific efforts and research needed toenhance an understanding of wetlands. At the sametime, this discussion is intended to contribute tohonest and frank debates among scientists, politicians,planners and the public, in order to reach a consensusfor the appropriate management of Brazilianwetlands. These discussions must include: (1) theestablishment of a programme and database for a

W. J. JUNK ET AL.18

Copyright # 2013 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 24: 5–22 (2014)

Page 15: Brazilian wetlands: their definition, delineation, and classi cation … · 2018-10-12 · Brazilian wetlands, their benefits to society, and the major threats endangering them.

Brazilian-wide wetland inventory following internationalstandards, e.g. the methodology used by the AsianWetland Inventory (AWI); (2) strengthening ofbasic and applied wetland research at selectedBrazilian universities and research institutions; (3)elaboration and implementation of a specificnational policy for the protection and sustainablemanagement of wetlands; (4) improvements in theefficiency of and cooperation among the manyorganizations working at different administrativelevels on wetlands and aquatic resources; (5)strengthening the cooperation between scientists,politicians, planners, and local stakeholders withrespect to wetland management and protection.Given the quickly changing economic, ecological,and social conditions in Brazil, and throughout theworld, and with the challenges that will be posed bythe anticipated dramatic changes in global climate,wetlands will play a crucial role that will benefitfuture generations.

REFERENCES

Adis J. 1997. Terrestrial invertebrates: survival strategies,group spectrum, dominance and activity patterns. In TheCentral Amazon Floodplain: Ecology of a Pulsing System,Junk WJ (ed.). Ecological Studies 126, Springer Verlag:Berlin/Heidelberg/New York; 299–317.

Adis J, Junk WJ. 2002. Terrestrial invertebrates inhabitinglowland river floodplains of Central Amazonia and CentralEurope: a review. Freshwater Biology 47: 711–731.

Araújo DSD, Henriques RPB. 1984. Análise florística dasrestingas do Estado do Rio de Janeiro. In Restingas.Origem, Processo e Estrutura, Lacerda LD, Araújo DSD,Cerqueira R, Turcq B (eds). CEUFF: Niterói, Brazil; 159–193.

Araújo GM, Barbosa AA, Arantes AA, Amaral AF. 2002.Composição florística de veredas no Município deUberlândia, MG. Revista Brasileira de Botânica 25: 475–493.

Araújo DSD, Scarano F, Sá CFC, Kurtz B, Zaluar HLT,Montezuma RCM, Oliveira RC. 1998. Comunidadesvegetais do Parque Nacional da Restinga de Jurubatiba. InEcologia das Lagoas Costeiras do Parque Nacional daRestinga de Jurubatiba e do Município de Macaé (RJ),Esteves FA (ed). NUPEM/UFRJ: Rio de Janeiro; 39–62.

Barletta M, Jaureguizar AJ, Baigun C, Fontoura NF,Agostinho AA, Almeida-Val VMF, Val AL, Torres RA,Jimenes-Segura LF, Giarrizzo T, et al. 2010. Fish andaquatic habitat conservation in South America: acontinental overview with emphasis on neotropical systems.Journal of Fish Biology 76: 2118–2176.

Beger M, Grantham HS, Pressey RL, Wilson KA, PetersonEL, Dorfman D, Mumby PJ, Lourival R, Brumbaugh DR,Possingham HP. 2010. Conservation planning forconnectivity across marine, freshwater, and terrestrialrealms. Biological Conservation 143: 565–575.

Bigarella JJ. 2001. Contribuição ao Estudo da PlanícieLitorânea do Estado do Paraná. Brazilian Archives ofBiology and Technology Jubilee Volume 2001: 65–110.

Bove CP, Gil ASB, Anjos RFB. 2003. Hidrófitasfanerogâmicas de ecossistemas aquáticos temporários daplanície costeira do Estado do Rio de Janeiro, Brasil. ActaBotanica Brasilica 17: 119–135.

BRASIL. 2012. National report on the implementation of theRamsar Convention on Wetlands. The Conference of theContracting Parties, Romania. (Available from: http://www.ramsar.org/pdf/cop11/nr/cop11-nr-brazil.pdf) [June 2012]

Brinson MM. 1993. A hydrogeomorphic classification forwetlands. Technical Report WRP-DE-4, U.S. ArmyEngineer Waterways Experiment Station, Vicksburg, MS.

BrinsonMM,Malvárez AI. 2002. Temperate freshwater wetlands:types, status, and threats. Environmental Conservation 29:115–133.

Casanova MT, Brock MA. 2000. How do depth, duration andfrequency of flooding influence the establishment of wetlandplant communities. Plant Ecology 147: 237–250.

Costa CSB, Davy AJ. 1992. Coastal saltmarsh communities ofLatin America. In Coastal Plant Communities of LatinAmerica, Seelinger U (ed). Academic Press: San Diego CA;179–199.

Costa AF, Dias ICA. 2001. Flora do Parque Nacional daRestinga de Jurubatiba e Arredores, Rio de Janeiro, Brasil:listagem florística e fitogeografia: Angiospermas, Pteridófitas,Algas Continentais. Editora do Museu Nacional: Rio deJaneiro.

Costanza R, d’Arge R, de Groot R, Farber S, Grasso M,Hannon B, Limburg K, Naeem S, O’Neill RV, Paruelo J,et al. 1997. The value of the world’s ecosystem services andnatural capital. Nature 387: 253–260.

Cowardin LM, Carter V, Golet FC, LaRoe ET. 1979.Classification of wetlands and deepwater habitats of theUnited States. US Department of the Interior, Fish andWildlife Service, Washington, DC.

Da Silva MP, Mauro R, Mourão G, Coutinho M. 2000.Distribuição e quantificação de classes de vegetação doPantanal através de levantamento aéreo. Revista Brasileirade Botânica 23: 143–152.

Darwall W, Smith K, Allen D, Seddon M, McGregor Reid G,Clausnitzer V, Kalkman V. 2008. Freshwater biodiversity – ahidden resource under threat. In The 2008 Review of theIUCN Red List of Threatened Species, Vié J-C, Hilton-TylorC, Stuard SN (eds). IUCN: Gland, Switzerland.

Davidson NC, Finlayson CM. 2007. Earth observation forwetland inventory, assessment and Monitoring. AquaticConservation: Marine and Freshwater Ecosystems 17: 219–228.

Diegues ACS. 1994. An Inventory of Brazilian Wetlands.IUCN – The World Conservation Union: Gland, Switzerland.

Diegues ACS. 2002. Povos e Águas. Núcleo de Apoio à Pesquisasobre Populações Humanas e Áreas Úmidas Brasileiras, 2nd edn.NUPAUB-USP (Universidade de Sao Paulo), São Paulo, Brazil.

Drechsler M, Lourival R, Possingham HP. 2009. Conservationplanning for successional landscapes. Ecological Modeling220: 438–450.

Eiten G. 1983. Classificação da vegetação do Brasil. CNPqBrasília: Brazil.

Erwin TL, Adis J. 1982. Amazonian inundation forests. Theirrole as short-term refuges and generators of species richnessand taxon pulses. In Biological Diversification in the

BRAZILIAN WETLANDS 19

Copyright # 2013 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 24: 5–22 (2014)

Page 16: Brazilian wetlands: their definition, delineation, and classi cation … · 2018-10-12 · Brazilian wetlands, their benefits to society, and the major threats endangering them.

Tropics, Prance GT (ed). Columbia University Press: NewYork; 358–371.

Esteves FA (ed). 2011: Fundamentos de Limnologia. Ed.Interciência: Rio de Janeiro.

Felfili JM. 1995. Diversity, structure and dynamics of a galleryforest in Central Brazil. Vegetatio 117: 1–15.

Finlayson CM. 2012. Forty years of wetland conservation andwise use. Aquatic Conservation: Marine and FreshwaterEcosystems 22: 139–143.

Finlayson CM, Van der Valk AG. 1995. Wetland classificationand inventory: a summary. Vegetatio 118: 185–192.

Gopal B, Sah M. 1995. Inventory and classification of wetlandsin India. Vegetatio 118: 39–48.

Gopal B, Junk WJ, Davis JA (eds). 2000. Biodiversity inWetlands: Assessment, Function and Conservation, Volume1. Backhuys Publishers: Leiden, The Netherlands.

Hamilton SK, Sippel SJ, Melack JM. 1996. Inundationpatterns in the Pantanal wetland of South Americadetermined from passive microwave remote sensing. Archivfür Hydrobiologie 137: 1–23.

Hamilton SK, Sippel SJ, Melack JM. 2004. Seasonalinundation patterns in two large savanna floodplains ofSouth América: the Llanos de Moxos (Bolivia) and theLlanos del Orinoco (Venezuela and Colombia).Hydrological Processes 18: 2103–2116.

Hoeinghaus DJ, Agostinho AA, Gomes LC, Pelicice FM,Okada EK, Latini JD, Kashiwaqui EAL, WinemillerKO. 2009. Effects of river impoundment on ecosystemservices of large tropical rivers: embodied energy andmarket value of artisanal fisheries. Conservation Biology23: 1222–1231.

Householder JE, Janovec JP, Page S. 2012. Peatlands of theMadre de Dios River of Peru: distribution, geomorphology,and habitat diversity. Wetlands 32: 359–368.

IPCC. 2007. Climate Change 2007: synthesis report. Part of theWorking Group III contribution to the fourth assessmentreport of the Intergovernmental Panel on Climate Change.Cambridge University Press: Cambridge.

Irgang BE, Gastal CVS. 1996. Macrófitas aquáticas da planíciecosteira do RS. CPG-Botanica, UFRGS (UniversidadeFederal de Rio Grande do Sul), Porto Alegre, Brazil.

Irgang BE, Pedralli G, Waechter JL. 1984. Macrófitasaquáticas da estação ecológica do Taim, Rio Grande doSul, Brasil. Rossléria 6: 395–405.

Junk WJ. 2000. Mechanisms for development and maintenanceof biodiversity in neotropical floodplains. In Biodiversity inWetlands: Assessment, Function and Conservation. Vol. 1,Gopal B, Junk WJ, Davis JA (eds). Backhuys Publishers:Leiden, The Netherlands; 119–139.

Junk WJ. 2005. Flood pulsing and the linkages betweenterrestrial, aquatic, and wetland systems. Proceedings of theInternational Association for Theoretical and AppliedLimnology 29: 11–38.

Junk WJ. 2007. Freshwater fishes of South America: theirbiodiversity, fisheries, and habitats: a synthesis. AquaticEcosystem Health & Management 10: 228–242.

Junk WJ, Piedade MTF. 1993. Herbaceous plants of theAmazon floodplain near Manaus: species diversity andadapations to the flood pulse. Amazoniana 12: 467–484.

Junk WJ, Piedade MTF. 1997. Plant life in the floodplain withspecial reference to herbaceous plants. In The CentralAmazon Floodplain: Ecology of a Pulsing System, Junk

WJ (ed). Ecological Studies, Vol 126. Springer Verlag:Berlin/Heidelberg/New York; 147–186.

Junk WJ, Piedade MTF, Schöngart J, Wittmann F. 2012b. Aclassification of major natural habitats of Amazonianwhitewater river floodplains (várzeas). Wetlands Ecologyand Management 20: 461–475.

Junk WJ, Sousa PT, Nunes da Cunha C, Piedade MTF,Candotti E. 2012a. Inundações catastróficas e deslizamentode barrancos em Minas Gerais e o novo Código florestal.Jornal da Ciência, Sociedade Brasileira para o Progresso daCiência. (Available from: http://www.jornaldaciencia.org.br/Detalhe.jsp?id=81006) (accessed on 20.09.2012).

Junk WJ, Wantzen KM. 2004. The flood pulse concept: newaspects, approaches, and applications – an update. InProceedings of the 2nd International Symposium on theManagement of Large Rivers for Fisheries, Volume 2,Welcome RL, Petr T (eds). Food and AgricultureOrganization & Mekong River Commission FAO RegionalOffice for Asia and the Pacific: Bangkok, Cambodia;117–149.

Junk WJ, Bayley PB, Sparks RE. 1989. The flood pulse conceptin river–floodplain systems. Canadian Special Publications forFisheries and Aquatic Sciences 106: 110–127.

Junk WJ, Nunes da Cunha C, Wantzen KM, Petermann P,Strüssmann C, Marques MI, Adis J. 2006. Biodiversity andits conservation in the Pantanal of Mato Grosso, Brazil.Aquatic Sciences 68: 278–309.

Junk WJ, Piedade MTF, Schöngart J, Cohn-Haft M,Adeney JM, Wittmann F. 2011. A classification ofmajor naturally-occurring Amazonian lowland wetlands.Wetlands 31: 623–640.

Kahn F. 1991. Palms as key swamp forest resources inAmazonia. Forest Ecology and Management 38: 133–142.

Kjerfve B, Lacerda LD. 1993. Mangroves of Brazil. InConservation and Sustainable Utilization of Mangrove Forestsin Latin America and Africa Regions. Vol.2 Part I- LatinAmerica, Lacerda LD (ed). Mangrove EcosystemsTechnical Reports ITTO/ISME Project PD114/90 (F):Okinawa, Japan; 245–272.

Lacerda LD. 2001. Mangrove Ecosystems: Function andManagement. Springer Verlag: Berlin.

Lacerda LD, Vaisman AG, Maia LP, Cunha E, Silva CAR.2006. Relative importance of nitrogen and phosphorusemissions from shrimp farming and other anthropogenicsources for six estuaries along the NE Brazilian coast.Aquaculture 253: 433–446.

Lopez A, Finlayson CM, Begg G, Davies J, Tagi K, Lowry J.2002. Asian Wetland Inventory (AWI) – a planning andmanagement tool for wetland conservation and wise usethrough multi-stakeholder involvement. In Developing aproposed framework for a wetland inventory, assessment andmonitoring system (WIAMS) in Malaysia, Murugadas(compiler). Proceedings of the workshop on Developing aproposed framework for a wetland inventory, assessment andmonitoring system (WIAMS) in Malaysia, April 2002, KualaLumpur, Malaysia, Wetlands International – MalaysiaProgramme, Petaling Jaya, Malaysia; 37–48.

Lourival R, Drechsler M, Watts ME, Game ET, PossinghamHP. 2011. Planning for reserve adequacy in dynamiclandscapes; maximizing future representation of vegetationcommunities under flood disturbance in the Pantanalwetland. Diversity and Distributions 17: 297–310.

W. J. JUNK ET AL.20

Copyright # 2013 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 24: 5–22 (2014)

Page 17: Brazilian wetlands: their definition, delineation, and classi cation … · 2018-10-12 · Brazilian wetlands, their benefits to society, and the major threats endangering them.

Maia LP, Lacerda LD,Monteiro LHU, Souza GM. 2006.Atlasdos Manguezais do Nordeste do Brasil: Avaliação das Áreas deManguezais dos Estados do Piauí, Ceará, Rio Grande doNorte, Paraíba e Pernambuco. SEMACE: Fortaleza.

Maltchik L, Costa ES, Becker CG, Oliveira AE. 2003.Inventory of wetlands of Rio Grande do Sul (Brazil).Pesquisas Botânica 53: 89–100.

Maltchik L, Costa MAJ, Duarte MCD. 1999. Inventory ofBrazilian semiarid shallow lakes. Anais da AcademiaBrasileira de Ciências 71: 801–808.

Maltchik L, Rolon AS, Guadagnini DL, Stenert C. 2004.Wetlands of Rio Grande do Sul, Brazil: a classification withemphasis on plant communities. Acta LimnologicaBrasiliensia 16: 137–151.

Melack JM, Hess LL. 2010. Remote sensing of the distributionand extent of wetlands in the Amazon basin. In AmazonFloodplain Forests: Ecophysiology, Biodiversity andSustainable Management, Junk WJ, Piedade MTF,Wittmann F, Schöngart J, Parolin P (eds). EcologicalStudies 210, Springer Verlag: Berlin/Heidelberg/New York;43–59.

Menezes MP, Berger U, Mehlig U. 2008. Mangrove vegetationin Amazonia: a review of studies from the coast of Pará andMaranhão States, north Brazil. Acta Amazonica 38: 403–420.

Millennium Ecosystem Assessment. 2005. Ecosystems andhuman wellbeing: wetlands and water. (Available from:http://www.unep.org/maweb/documents/document.358.aspx.pdf) (accessed on: 20.09.2012).

Mitsch WJ, Day JW. 2006. Restoration of wetlands in theMississippi–Ohio–Missouri (MOM) River Basin: experienceand needed research. Ecological Engineering 26: 55–69.

Mitsch WJ, Gosselink JG. 2008. Wetlands. John Wiley:Hoboken, NJ.

Naiman RJ, Decamps H. 1997. The ecology of interfaces:riparian zones. Annual Review of Ecology and Systematics28: 621–658.

Neiff JJ. 2001. Humedales de la Argentina: sinopsis, problemasy perspectivas futuras. In Funciones de los humedales, calidadde vida y agua segura, Cirelli AF (ed.). El agua enIberoamérica, Publ. CYTED: Buenos Aires; 83–112.

Nunes da Cunha C, Junk WJ. 2011a. A preliminaryclassification of habitats of the Pantanal of Mato Grossoand Mato Grosso do Sul, and its relation to national andinternational classification systems. In The Pantanal:Ecology, Biodiversity and Sustainable Management of aLarge Neotropical Seasonal Wetland, Junk WJ, da Silva CJ,Nunes da Cunha C, Wantzen KM (eds). Pensoft: Sofia/Moscow;127–142.

Nunes da Cunha C, Junk WJ. 2011b. Landscape units of thePantanal: structure, function and human use. In ThePantanal: Ecology, Biodiversity and SustainableManagement of a Large Neotropical Seasonal Wetland,Junk WJ, da Silva CJ, Nunes da Cunha C, Wantzen KM(eds). Pensoft: Sofia/Moscow; 299–324.

Nunes da Cunha C, Junk WJ, Leitão-Filho H. 2007. Woodyvegetation in the Pantanal of Mato Grosso, Brazil: apreliminary typology. Amazoniana 19: 159–184.

Piedade MTF, Junk WJ, Long SP. 1991. The productivity ofthe C4 grass Echinochloa polystachia on the Amazonfloodplain. Ecology 72: 1456–1463.

Piedade MTF, Junk WJ, Sousa PT, Nunes da Cunha C,Schöngart J, Wittmann F, Candotti E, Girard P. 2012. As

áreas úmidas no âmbito do Código Florestal brasileiro. InCódigo Florestal e a ciência: o que nossos legisladores aindaprecisam saber. Sumários executivos de estudos científicossobre impactos do projeto de Código Florestal, Comitê Brasilem Defesa das Florestas e do Desenvolvimento Sustentável(ed). Comitê Brasil: Brasília, Brazil; 9–17.

Pott VJ, Pott A. 2000. Plantas Aquáticas do Pantanal.EMBRAPA: Brasília, Brazil.

Ribeiro JF, Walter BMT. 1998. Fitofisionomias do biomacerrado. In Cerrado: ambiente e flora, Sano SM, AlmeidaSP (eds). EMBRAPA-CPAC: Planaltina; 89–166.

Ribeiro MC, Metzger JP, Martensen AC, Ponzoni FJ, HirotaMM. 2009. The Brazilian Atlantic Forest: how much is left,and how is the remaining forest distributed? Implicationsfor conservation. Biological Conservation 142: 1141–1153.

Rolon AS, Homem HF, Maltchik L. 2010. Aquaticmacrophytes in natural and managed wetlands of RioGrande do Sul State, Southern Brazil. Acta LimnologicaBrasiliensia 22: 133–146.

Ruokolainen K, Schulman L, Tuomisto H. 2001. OnAmazonian peatlands. International Mire ConservationGroup Newsletter 4: 8–10.

Salati E, Marques J. 1984. Climatology of the Amazon region.In The Amazon - Limnology and Landscape Ecology of aMighty Tropical River and its Basin, Sioli H (ed.). Dr W.Junk Publishers: Dordrecht, The Netherlands; 85–126.

Scarano FR. 2006. Plant community structure and function in aswamp forest within the Atlantic rain forest complex: asynthesis. Rodriguésia 57: 491–502.

SCBD. 2010. Global Biodiversity Outlook 3. Secretariat of theConvention on Biodiversity: Montreal, Canada.

Schaeffer-Novelli Y, Cintrón-Molero G, Adaime RR,Camargo TM. 1990. Variability of mangrove ecosystemsalong the Brazilian coast. Estuaries 13: 204–218.

Schöngart J, Wittmann F, Worbes M. 2010. Biomass and netprimary production of Central Amazonian floodplainforests. In Amazonian Floodplain Forests: Ecophysiology,Biodiversity and Sustainable Management, Junk WJ,Piedade MTF, Wittmann F, Schöngart J, Parolin P (eds).Ecological Studies 210, Springer Verlag: Berlin/ Heidelberg/NewYork; 347–388.

Scott DA, Jones TA. 1995. Classification and inventory ofwetlands: a global overview. Vegetatio 118: 3–16.

Seidl AF, Moraes AS. 2000. Global valuation of ecosystemservices: application to the Pantanal da Nhecolandia,Brazil. Ecological Economics 33: 1–6.

Semeniuk CA, Semeniuk V. 1995. A geomorphic approach toglobal classification for inland wetlands. Vegetatio 118:103–124.

TEEB. 2013. TEEB for Water and Wetlands. Institute forEuropean Environmental Policy (IEEP) & Ramsar Secretariat,London and Brussels. (Available from: http://www.teebtest.org/wp-content/uploads/2013/04/TEEB_WaterWetlands_Report_2013.pdf) (accessed on: 15.04.2013).

Thomaz SM, Agostinho AA, Hahn NS. 2004. The UpperParaná River and its Floodplain: Physical Aspects, Ecologyand Conservation. Backhuys Publishers: Leiden, TheNetherlands.

Wantzen KM. 2003. Cerrado streams – characteristics of athreatened freshwater ecosystem type on the TertiaryShields of Central South America. Amazoniana 17:481–502.

BRAZILIAN WETLANDS 21

Copyright # 2013 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 24: 5–22 (2014)

Page 18: Brazilian wetlands: their definition, delineation, and classi cation … · 2018-10-12 · Brazilian wetlands, their benefits to society, and the major threats endangering them.

Wittmann F. 2012. Tree species composition and diversity inBrazilian freshwater floodplains. In Mycorrhiza: Occurrencein Natural and Restored Environments, Pagano MC (ed.).Nova Science Publishers: New York; 223–263.

Wittmann F, Householder E, Piedade MTF, Assis RL,Schöngart J, Parolin P, Junk WJ. 2013. Habitat specificity,endemism and the neotropical distribution of Amazonianwhite-water floodplain trees. Ecography 36: 690–707.

Wittmann F, Schöngart J, Junk WJ. 2010. Phytogeography,species diversity, community structure and dynamics ofcentral Amazonian floodplain forests. In AmazonianFloodplain Forests: Ecophysiology, Biodiversity andSustainable Management, Junk WJ, Piedade MTF,Wittmann F, Schöngart J, Parolin P (eds). EcologicalStudies 210, Springer Verlag: Berlin/Heidelberg/New York;61–102.

W. J. JUNK ET AL.22

Copyright # 2013 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 24: 5–22 (2014)